Existence and stability of standing pulses in neural networks

This dissertation studies a one dimensional neural network rate model that supports localized self-sustained solutions. These solutions could be an analog for working memory in the brain. Working memory refers to the temporary storage of information necessary for performing different mental tasks. Cortical neurons that show persistent activity are observed in animals during working memory tasks. The physical process underlying this persistent activity could be due to self-sustained network activity of the neurons in the brain.The term `bump' has been coined to imply a spatially localized persistent activity state that is sustained internally by a network of neurons. Many researchers have analyzed the bump state using Firing rate models with either the Heaviside gain function or a saturating sigmoidal one. These gain functions imply that neurons begin to fire once their synaptic input reaches threshold, and the firing rate saturates to a maximal value almost immediately. However, cortical neurons that exhibit persistent activity usually are well below their maximal attainable rate. To resolve this paradox, I study a single population rate model using a biophysically relevant firing rate function.I consider the existence and the stability of standing single-pulse solutions of an integro-diferential neural network equation. In this network, the synaptic coupling has local excitatory coupling with distal lateral inhibition and the non-saturating gain function is piece-wise linear. A standing pulse solution of this network is a synaptic input pattern that supports a bump state. I show that the existence condition for single-pulses of the integro-differential equation can be reducedto the solution of an algebraic system. With this condition, I map out the shape of the pulsesfor different coupling weights and gains. By a similar approach, I also find the conditions for the existences of dimple-pulses and double-pulses. For a fixed gain and connectivity, there are at least two single-pulse solutions - a "large" one and a "small" one. However, more than two single-pulses can coexist depending on the parameter range. To have standing single-pulses, the gain function and synaptic coupling are both important.I also derive a stability criteria for the standing pulse solutions. I show that the large pulse is stable and the small pulse is unstable. If there are more than two pulse solutions coexisting, the first pulse is the small one and it is unstable. The second one is a large stable pulse. The third pulse is wider than the second one and it is unstable. More importantly, the second single-pulse (which could be a dimple pulse) is bistable with the "all-off" state. The stable pulse represents the memory. When the network is switched to the "all-off" state, the memory is erased.

[1]  Todd Kapitula,et al.  The Evans function for nonlocal equations , 2004 .

[2]  B. Richmond,et al.  Intrinsic dynamics in neuronal networks. II. experiment. , 2000, Journal of neurophysiology.

[3]  P. Goldman-Rakic,et al.  Correlated discharges among putative pyramidal neurons and interneurons in the primate prefrontal cortex. , 2002, Journal of neurophysiology.

[4]  H. Sompolinsky,et al.  Theory of orientation tuning in visual cortex. , 1995, Proceedings of the National Academy of Sciences of the United States of America.

[5]  H. Sompolinsky,et al.  Mexican hats and pinwheels in visual cortex , 2003, Proceedings of the National Academy of Sciences of the United States of America.

[6]  Richard K Miller Introduction to Differential Equations , 1982 .

[7]  Stephen Coombes,et al.  Evans Functions for Integral Neural Field Equations with Heaviside Firing Rate Function , 2004, SIAM J. Appl. Dyn. Syst..

[8]  S. Amari,et al.  Existence and stability of local excitations in homogeneous neural fields , 1979, Journal of mathematical biology.

[9]  Linghai Zhang,et al.  On stability of traveling wave solutions in synaptically coupled neuronal networks , 2003, Differential and Integral Equations.

[10]  P. Goldman-Rakic,et al.  Division of labor among distinct subtypes of inhibitory neurons in a cortical microcircuit of working memory. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[11]  R. A. Davidoff From Neuron to Brain , 1977, Neurology.

[12]  Alan R. Champneys,et al.  On solitary waves of a piecewise linear suspended beam model , 1997 .

[13]  Carson C. Chow,et al.  Localized Bumps of Activity Sustained by Inhibition in a Two-Layer Thalamic Network , 2004, Journal of Computational Neuroscience.

[14]  L. Delves,et al.  Computational methods for integral equations: Frontmatter , 1985 .

[15]  John Evans,et al.  Nerve Axon Equations: II Stability at Rest , 1972 .

[16]  X. Wang,et al.  Synaptic Basis of Cortical Persistent Activity: the Importance of NMDA Receptors to Working Memory , 1999, The Journal of Neuroscience.

[17]  Boris S. Gutkin,et al.  Multiple Bumps in a Neuronal Model of Working Memory , 2002, SIAM J. Appl. Math..

[18]  Masayasu Mimura,et al.  Layer oscillations in reaction-diffusion systems , 1989 .

[19]  C. Laing,et al.  Two-bump solutions of Amari-type models of neuronal pattern formation , 2003 .

[20]  Jonathan E. Rubin,et al.  Sustained Spatial Patterns of Activity in Neuronal Populations without Recurrent Excitation , 2004, SIAM J. Appl. Math..

[21]  G. Lord,et al.  Waves and bumps in neuronal networks with axo-dendritic synaptic interactions , 2003 .

[22]  Charalambos D. Aliprantis,et al.  Principles of Real Analysis , 1981 .

[23]  N. Rashevsky,et al.  Mathematical biology , 1961, Connecticut medicine.

[24]  Stephen Wolfram,et al.  The Mathematica Book , 1996 .

[25]  Bard Ermentrout,et al.  Spatially Structured Activity in Synaptically Coupled Neuronal Networks: I. Traveling Fronts and Pulses , 2001, SIAM J. Appl. Math..

[26]  Bard Ermentrout,et al.  Spatially Structured Activity in Synaptically Coupled Neuronal Networks: II. Lateral Inhibition and Standing Pulses , 2001, SIAM J. Appl. Math..

[27]  P. Goldman-Rakic,et al.  Temporally irregular mnemonic persistent activity in prefrontal neurons of monkeys during a delayed response task. , 2003, Journal of neurophysiology.

[28]  Bard Ermentrout,et al.  Reduction of Conductance-Based Models with Slow Synapses to Neural Nets , 1994, Neural Computation.

[29]  J. Fuster The Prefrontal Cortex , 1997 .

[30]  S Grossberg,et al.  Some developmental and attentional biases in the contrast enhancement and short term memory of recurrent neural networks. , 1975, Journal of theoretical biology.

[31]  H. Eom Green’s Functions: Applications , 2004 .

[32]  K. Atkinson The Numerical Solution of Integral Equations of the Second Kind , 1997 .

[33]  L. Peletier,et al.  Spatial Patterns: Higher Order Models in Physics and Mechanics , 2001 .

[34]  J. Deuchars,et al.  Temporal and spatial properties of local circuits in neocortex , 1994, Trends in Neurosciences.

[35]  W. Rudin Principles of mathematical analysis , 1964 .

[36]  Carlo R. Laing,et al.  PDE Methods for Nonlocal Models , 2003, SIAM J. Appl. Dyn. Syst..

[37]  E. Zeidler Nonlinear functional analysis and its applications , 1988 .

[38]  Tosio Kato Perturbation theory for linear operators , 1966 .

[39]  J. Cowan,et al.  Excitatory and inhibitory interactions in localized populations of model neurons. , 1972, Biophysical journal.

[40]  P. Goldman-Rakic,et al.  Mnemonic coding of visual space in the monkey's dorsolateral prefrontal cortex. , 1989, Journal of neurophysiology.

[41]  Jonathan E. Rubin,et al.  A NONLOCAL EIGENVALUE PROBLEM FOR THE STABILITY OF A TRAVELING WAVE IN A NEURONAL MEDIUM , 2004 .

[42]  George Weiss,et al.  Integral Equation Methods , 1969 .

[43]  Steven H. Strogatz,et al.  Nonlinear Dynamics and Chaos , 2024 .

[44]  S. Wiggins Introduction to Applied Nonlinear Dynamical Systems and Chaos , 1989 .

[45]  Matiur Rahman,et al.  Complex Variables and Transform Calculus , 1997 .

[46]  E. Kreyszig Introductory Functional Analysis With Applications , 1978 .

[47]  S. Amari Dynamics of pattern formation in lateral-inhibition type neural fields , 1977, Biological Cybernetics.

[48]  Carson C. Chow,et al.  Existence and Stability of Standing Pulses in Neural Networks: II. Stability , 2004, SIAM J. Appl. Dyn. Syst..

[49]  B. Richmond,et al.  Intrinsic dynamics in neuronal networks. I. Theory. , 2000, Journal of neurophysiology.

[50]  Bard Ermentrout,et al.  Simulating, analyzing, and animating dynamical systems - a guide to XPPAUT for researchers and students , 2002, Software, environments, tools.

[51]  A. Balakrishnan Applied Functional Analysis , 1976 .

[52]  I. N. Sneddon,et al.  Boundary value problems , 2007 .

[53]  Boris S. Gutkin,et al.  Turning On and Off with Excitation: The Role of Spike-Timing Asynchrony and Synchrony in Sustained Neural Activity , 2001, Journal of Computational Neuroscience.

[54]  Eva Part-Enander,et al.  The Matlab Handbook , 1996 .

[55]  H. Spinnler The prefrontal cortex, Anatomy, physiology, and neuropsychology of the frontal lobe, J.M. Fuster. Raven Press, New York (1980), IX-222 pages , 1981 .

[56]  G. Folland Fourier analysis and its applications , 1992 .

[57]  V. G. Yakhno,et al.  Generation of Collective-Activity Structures in a Homogeneous Neuron-Like Medium i: Bifurcation Analysis of Static Structures , 1996 .

[58]  B. Ermentrout Neural networks as spatio-temporal pattern-forming systems , 1998 .

[59]  S. Grossberg,et al.  Pattern formation, contrast control, and oscillations in the short term memory of shunting on-center off-surround networks , 1975, Biological Cybernetics.

[60]  L. Abbott,et al.  A model of multiplicative neural responses in parietal cortex. , 1996, Proceedings of the National Academy of Sciences of the United States of America.

[61]  Y. Kuznetsov Elements of Applied Bifurcation Theory , 2023, Applied Mathematical Sciences.

[62]  G. E. Alexander,et al.  Neuron Activity Related to Short-Term Memory , 1971, Science.

[63]  Frank G. Garvan,et al.  The MAPLE Book , 2001 .

[64]  B W Connors,et al.  Intrinsic neuronal physiology and the functions, dysfunctions and development of neocortex. , 1994, Progress in brain research.

[65]  John Evans Nerve Axon Equations: III Stability of the Nerve Impulse , 1972 .

[66]  Xiao-Jing Wang,et al.  A Model of Visuospatial Working Memory in Prefrontal Cortex: Recurrent Network and Cellular Bistability , 1998, Journal of Computational Neuroscience.

[67]  R. Desimone,et al.  Neural Mechanisms of Visual Working Memory in Prefrontal Cortex of the Macaque , 1996, The Journal of Neuroscience.

[68]  P. Goldman-Rakic,et al.  Coding Specificity in Cortical Microcircuits: A Multiple-Electrode Analysis of Primate Prefrontal Cortex , 2001, The Journal of Neuroscience.

[69]  D. Griffel Applied functional analysis , 1982 .

[70]  P. Goldman-Rakic Cellular basis of working memory , 1995, Neuron.

[71]  Norman Morrison,et al.  Introduction to Fourier Analysis , 1994, An Invitation to Modern Number Theory.

[72]  Michael A. Arbib,et al.  The handbook of brain theory and neural networks , 1995, A Bradford book.

[73]  Carson C. Chow,et al.  Stationary Bumps in Networks of Spiking Neurons , 2001, Neural Computation.

[74]  J. Cowan,et al.  A mathematical theory of the functional dynamics of cortical and thalamic nervous tissue , 1973, Kybernetik.

[75]  M. Goldberg,et al.  Oculocentric spatial representation in parietal cortex. , 1995, Cerebral cortex.

[76]  H S Seung,et al.  How the brain keeps the eyes still. , 1996, Proceedings of the National Academy of Sciences of the United States of America.

[77]  Xiao-Jing Wang Synaptic reverberation underlying mnemonic persistent activity , 2001, Trends in Neurosciences.

[78]  P. Zweifel Advanced Mathematical Methods for Scientists and Engineers , 1980 .